insectInsecta or Hexapodaany member of the largest class of the phylum Arthropoda, which is itself the largest of the animal phyla. Insects have segmented bodies, jointed legs, and external skeletons (exoskeletons). Insects are distinguished from other arthropods by their body, which is divided into three major regions: (1) the head, which bears the mouthparts, eyes, and a pair of antennae, (2) the three-segmented thorax, which usually has three pairs of legs (hence “Hexapoda”) in adults and usually one or two pairs of wings, and (3) the many-segmented abdomen, which contains the digestive, excretory, and reproductive organs.

In a popular sense, “insect” usually refers to familiar pests or disease carriers, such as bedbugs, houseflies, clothes moths, Japanese beetles, aphids, mosquitoes, fleas, horseflies, and hornets, or to conspicuous groups, such as butterflies, moths, and beetles. Many insects, however, are beneficial from a human viewpoint; they pollinate plants, produce useful substances, control pest insects, act as scavengers, and serve as food for other animals (see below Importance). Furthermore, insects are valuable objects of study in elucidating many aspects of biology and ecology. Much of our knowledge of genetics has been gained from fruit fly experiments and of population biology from flour beetle studies. Insects are often used in investigations of hormonal action, nerve and sense organ function, and many other physiological processes. Insects are also used as environmental quality indicators to assess water quality and soil contamination and are the basis of many studies of biodiversity.

General features

In numbers of species and individuals and in adaptability and wide distribution, insects are perhaps the most eminently successful group of all animals. They dominate the present-day land fauna with about 1,000,000 described species. This represents about three-fourths of all described animal species. Entomologists estimate the actual number of living insect species could be as high as 5,000,000 to 10,000,000. The orders that contain the greatest numbers of species are Coleoptera (beetles), Lepidoptera (butterflies and moths), Hymenoptera (ants, bees, wasps), and Diptera (true flies).

Appearance and habits

The majority of insects are small, usually less than 6 mm (0.2 inch) long, although the range in size is wide. Some of the feather-winged beetles and parasitic wasps are almost microscopic, while some tropical forms, such as the hercules beetles, African goliath beetles, certain Australian stick insects, and some Asian and South American moths, can be as large as 16 cm (6.3 inches).

In many species the difference in body structure between the sexes is pronounced, and knowledge of one sex may give few clues to the appearance of the other sex. In some, such as the twisted-wing insects (Strepsiptera), the female is a mere inactive bag of eggs, and the winged male is one of the most active insects known. Modes of reproduction are quite diverse, and reproductive capacity is generally high. Some insects, such as the mayflies, feed only in the immature or larval stage and go without food during an extremely short adult life. Among social insects, queen termites may live for up to 50 years, whereas some adult mayflies live less than two hours.

Some insects advertise their presence to the other sex by flashing lights, and many imitate other insects in colour and form and thus avoid or minimize attack by predators that feed by day and find their prey visually, as do birds, lizards, and other insects.

Behaviour is diverse, from the almost inert parasitic forms, whose larvae lie in the nutrient bloodstreams of their hosts and feed by absorption, to dragonflies that pursue victims in the air, tiger beetles that outrun prey on land, and dytiscid beetles that outswim prey in water.

In some cases the adult insects make elaborate preparations for the young, in others the mother alone defends or feeds her young, and in still others the young are supported by complex insect societies. Some colonies of social insects, such as tropical termites and ants, may reach populations of millions of inhabitants.

Distribution and abundance

No scientist familiar with insects has attempted to estimate individual numbers beyond areas of a few acres or a few square miles in extent. Figures soon become so large as to be incomprehensible. The large populations and great variety of insects are related to their small size, high rates of reproduction, and abundance of suitable food supplies. Insects abound in the tropics, both in numbers of different kinds and in numbers of individuals.

If the insects (including the young and adults of all forms) are counted on a square yard (0.84 m2) of rich moist surface soil, 500 are found easily and 2,000 are not unusual in soil samples in the north temperate zone. This amounts to roughly 4,000,000 insects on one moist acre (0.41 ha). In such an area only an occasional butterfly, bumblebee, or large beetle, supergiants among insects, probably would be noticed. Only a few thousand species, those that attack man’s crops, herds, and products and those that carry disease, interfere with human life seriously enough to require control measures.

Insects are adapted to every land and freshwater habitat where food is available, from deserts to jungles, from glacial fields and cold mountain streams to stagnant, lowland ponds and hot springs. Many live in brackish water up to 110 the salinity of seawater, a few live on the surface of seawater, and some fly larvae can live in pools of crude petroleum, where they eat other insects that fall in.

Importance

Role in nature

Insects play many important roles in nature. They aid bacteria, fungi, and other organisms in the decomposition of organic matter and in soil formation. The decay of carrion, for example, brought about mainly by bacteria, is accelerated by the maggots of flesh flies and blowflies. The activities of these larvae, which distribute and consume bacteria, are followed by those of moths and beetles, which break down hair and feathers. Insects and flowers have evolved together. Many plants depend on insects for pollination. Some insects are predators of others.

Commercial significance

Certain insects provide sources of commercially important products such as honey, silk, wax, dyes, or pigments, all of which can be of direct benefit to man. Because they feed on many types of organic matter, insects can cause considerable agricultural damage. Insect pests devour crops of food or timber, either in the field or in storage, and convey infective micro-organisms to crops, farm animals, and human beings. The technology for combatting such pests constitutes the applied sciences of agricultural and forest entomology, stored product entomology, medical and veterinary entomology, and urban entomology.

Insects as a source of raw materials

For primitive peoples who gathered food, insects were a significant food source. Grasshopper plagues, termite swarms, large palm weevil grubs, and other insects are still sources of protein in some countries. The dry scaly excreta of coccids (Homoptera) on tamarisk or larch trees is the source of manna in the Sinai Desert. Coccids were once the source of the crimson dye kermes. The cochineal, or carmine, from Dactylopius scale insects found on Mexican cacti, was used for dying cloth by the Aztecs and is used today as a dye in foods, makeup, drugs, and textiles. Several insect waxes are used commercially, especially beeswax and lac wax. The resinous product of the lac insect Tachardia (Homoptera), which is cultured for this purpose, is the source of commercial shellac.

Two of the most important domesticated insects are the silkworm (Lepidoptera) and the honeybee (Hymenoptera). Some coarse silks are produced from the cocoons of large wild silkworm species. Most commercial silks, however, come from the silkworm Bombyx mori. This insect is unknown in the wild state and exists only in culture. It was domesticated in China thousands of years ago, and selective breeding, notably in China and Japan, has produced many specialized strains. The honeybee is a close relative of existing wild bees. In the Middle Ages, honey was Europe’s most important sweetener, and both beeswax and honey are still articles of commerce. However, the major importance of honeybees lies in their pollination of fruit trees and other crops.

Insect damage to commercial products

When insects that break down dead trees invade structural timbers in buildings, they become pests. This is true of insects such as dermestid beetles and various tineid moths that ecologically are latecomers to carcasses and are capable of breaking down the keratin in hair and feathers. When these insects invade skins, furs, and wool garments or carpets, they can become problems for humans.

In many hot, dry climates, as in North Africa or the plains of India, ripened grain in the fields is invaded by certain beetles and moths. When the grain is harvested, these insects thrive in the grain stores. They can be carried throughout the world in commerce and have become universal pests of stored grain, dried fruit, tobacco, and other products. Quarantine and disinfestation methods are used to control importation of such insects from grain-exporting countries.

Agricultural significance

Ecological factors

Many insects are plant feeders, and when the plants are of agricultural importance, man is often forced to compete with these insects. Populations of insects are limited by such factors as unfavourable weather, predators and parasites, and viral, bacterial, and fungal diseases, as well as many other factors that operate to make insect populations stable. Agricultural methods that encourage the planting of ever larger areas to single crops, which provides virtually unlimited food resources, has removed some of these regulating factors and allowed the rate of population growth of insects that attack those crops to increase. This increases the probability of great infestations of certain insect pests. Many natural forests, which form similar giant monocultures, always seem to have been subject to periodic outbreaks of destructive insects.

In some agricultural monocultures, nonnative insect pests have been accidentally introduced along with a crop, but without also bringing along its full range of natural enemies. This has occurred in the United States with the oyster scale (Lecanium) of apple, the cottony cushion scale (Icerya) of citrus, the European corn borer (Pyrausta), and others. The Colorado potato beetle (Leptinotarsa), which caused appalling destruction a century ago, was a native insect of semidesert country. The beetle, which fed on the buffalo burr plant, adapted itself to a newly introduced and abundant diet of potatoes and thus escaped from all previous controlling factors. Similar situations often have been controlled by determining the major predators or parasites of an alien insect pest in its country of origin and introducing them as control agents. A classic example is the cottony cushion scale, which threatened the California citrus industry in 1886. A predatory ladybird beetle, the vedalia beetle (Rodolia cardinalis), was introduced from Australia, and within a year or two the scale insect had virtually disappeared. The success was repeated in every country where the scale insect had become established without its predators. In eastern Canada in in the early 1940s the European spruce sawfly (Gilpinia), which had caused immense damage, was completely controlled by the spontaneous appearance of a viral disease, perhaps unknowingly introduced from Europe. This event led to increased interest in using insect diseases as potential means of managing pest populations.

Damage to growing crops

Insects are responsible for two major kinds of damage to growing crops. First is direct injury done to the plant by the feeding insect, which eats leaves or burrows in stems, fruit, or roots. There are hundreds of pest species of this type, both in larvae and adults, among orthopterans, homopterans, heteropterans, coleopterans, lepidopterans, and dipterans. The second type is indirect damage in which the insect itself does little or no harm but transmits a bacterial, viral, or fungal infection into a crop. Examples include the viral diseases of sugar beets and potatoes, carried from plant to plant by aphids.

Although most insects grow and multiply in the crop they damage, certain grasshoppers are well-known exceptions. They can exist in a relatively harmless solitary phase for a number of years, during which time their numbers may increase. They then enter a gregarious phase, forming gigantic migratory swarms, which are transported by winds or flight for hundreds or thousands of miles. These swarms may completely destroy crops in an invaded region. The desert locust (Schistocerca gregaria) and migratory locust (Locusta migratoria) are two examples of this type of life cycle.

Medical significance

Insect damage to man and livestock also may be direct or indirect. Direct injury to man by insect stings and bites is of relatively minor importance, although swarms of biting flies and mosquitoes often make life almost intolerable, as do biting midges (sand flies) and salt-marsh mosquitoes. Persistent irritation by biting flies can cause deterioration in the health of cattle. Some blowflies, in addition to depositing their eggs in carcasses, also invade the tissue of living animals and man, a condition known as myiasis. An example of an insect that causes this condition is the screwworm fly (Cochliomyia) of the southern U.S. and Central America. In many parts of the world various blowflies infest the fleece and skin of living sheep. This infestation, called sheep-strike, causes severe economic damage.

Many major diseases of man are produced by micro-organisms conveyed by insects, which serve as vectors of pathogens. Malaria is caused by the protozoan Plasmodium, which spends part of its developmental cycle in Anopheles mosquitoes. Epidemic relapsing fever, caused by spirochetes, is transmitted to man by the human louse Pediculus. Leishmaniasis, caused by the protozoan Leishmania, is carried by the sand fly Phlebotomus. Sleeping sickness in man and a group of cattle diseases that are widespread in Africa and known as nagana are caused by protozoan trypanosomes transmitted by the bites of tsetse flies (Glossina). Under nonsanitary conditions the common housefly Musca can play an incidental role in the spread of human intestinal infections (e.g., typhoid, bacillary and amebic dysentery) by contamination of human food. The tularemia bacillus can be spread by deer fly bites, the bubonic plague bacillus by fleas, and the epidemic typhus rickettsia by the louse Pediculus. Various mosquitoes spread viral diseases (e.g., several encephalitis diseases; dengue and yellow fever in man and other animals).

The relationships among the various organisms are complex. Malaria, for example, has a different epidemiology in almost every country in which it occurs, with different Anopheles species responsible for its spread. These same complexities affect the spread of sleeping sickness. The relationships between man and some diseases are indirect. Plague, a disease of rodents transmitted by flea bites, is dangerous to man only when heavy mortality among domestic rats forces their infected fleas to attack man, thereby causing an outbreak of plague. Typhus, tularemia, encephalitis, and yellow fever also are maintained in animal reservoirs and spread occasionally to man.

Control of insect damage

The historical objective of the entomologist was primarily to develop and introduce modifications into the environment in such ways that diseases will not be spread by insects, and crops will not be damaged by them. This objective has been achieved in numerous cases. For example, in many cities flies no longer play a major role in spreading intestinal infections, and improved land drainage, housing of man and animals, and insecticide use has eliminated malaria in many parts of the world.

Massive outbreaks of the Colorado potato beetle in the 1860s led to the first large-scale use of insecticides in agriculture. These highly poisonous chemicals (e.g., Paris green, lead arsenate, concentrated nicotine) were used in large quantities. The continued search for effective synthetic compounds led in the early 1940s to the production of DDT, a remarkable compound that is highly toxic to most insects, nontoxic to man in small quantities (although cumulative effects may be severe), and long lasting in effect. Widely used in agriculture for many years, DDT was not the perfect insecticide. It often killed parasites as effectively as the pests themselves, creating ecological imbalances that permitted new pests to develop large populations. Furthermore, resistant strains of pests appeared. The environmental longevity of many early insecticides was also found to cause significant ecological problems. Similar difficulties were encountered with many successors to DDT, such as Dieldrin and Endrin.

In the course of developing effective insecticides, the primary emphases have been to reduce their potential to cause human health problems and their impact on the environment. Biological methods of pest management have become increasingly important as the use of undesirable insecticides decreases. Biological methods include introducing pest strains that carry lethal genes, flooding an area with sterile males (as was successfully done for the control of the screwworm fly), or developing new kinds of insecticide based on modifications of insects’ growth hormones. The sugar industry in Hawaii and the California citrus industry rely on biological control methods. Although these methods are not consistently effective, they are considered to be less harmful to the environment than are some chemicals.

Natural history

Life cycle

Egg

Most insects begin their lives as fertilized eggs. The chorion, or eggshell, is commonly pierced by respiratory openings that lead to an air-filled meshwork inside the shell. For some insects (e.g., cockroaches and mantids) a batch of eggs is cemented together to form an egg packet or ootheca. Insects may pass unfavourable seasons in the egg stage. Eggs of the springtail Sminthurus (Collembola) and of some grasshoppers (Orthoptera) pass summer droughts in a dry shrivelled state and resume development when moistened. Most eggs, however, retain their water although they may pass the winter in a state of arrested development, or diapause, usually at some early stage in embryonic development. However, dried eggs of Aedes mosquitoes enter a state of dormancy after development is complete and quickly hatch when placed in water.

The hatching of young larvae is achieved in several ways. Some, such as caterpillars, bite their way out of the egg. Many, such as the flea, have hatching spines with which they cut a slit in the shell. Some insect eggs have a preformed “escape cap” that the larva pops from the shell by increasing the pressure inside the egg. Depending on the species, this may be accomplished either by swallowing air and then constricting muscles in the body to exert pressure on the cap or by having an expandable region on the head (many Diptera have a ptilinum) that can be extended by hydraulic (blood) pressure. After hatching, the larva continues to distend itself in this way, although the ptilinum collapses back into the body, until the cuticle hardens.

Once formed, the insect cuticle cannot grow. Growth can occur only by a series of molts (ecdyses) during which new and larger cuticles form and old cuticles are shed. Molting makes possible large changes in body form.

Types of metamorphosis

In the most primitive wingless insects (apterygotes) such as the silverfish Lepisma, there is almost no change in form throughout growth to the adult. These are known as ametabolous insects. Among insects such as grasshoppers (Orthoptera), true bugs (Heteroptera), and homopterans (e.g., aphids, scale insects), the general form is constant until the final molt, when the larva undergoes substantial changes in body form to become a winged adult with fully developed genitalia. These insects, termed hemimetabolous, are said to undergo incomplete metamorphosis. The higher orders of insects, including Lepidoptera (butterflies and moths), Coleoptera (beetles), Hymenoptera (ants, wasps, and bees), Diptera (true flies), and several others, are termed holometabolous because larvae are totally unlike adults. These larvae undergo a series of molts with little change in form before they enter into complete metamorphosis, which includes molting first into pupae and then into fully winged adults.

Types of larvae

Larvae, which vary considerably in shape, are classified in five forms: eruciform (caterpillar-like), scarabaeiform (grublike), campodeiform (elongated, flattened, and active), elateriform (wireworm-like), and vermiform (maggot-like). The three types of pupae are: obtect, with appendages more or less glued to the body; exarate, with the appendages free and not glued to the body; and coarctate, which is essentially exarate but remaining covered by the cast skins (exuviae) of the next to the last larval instar (name given to the form of an insect between molts).

Role of hormones

Both molting and metamorphosis are controlled by hormones. Molting is initiated when sensory receptors in the body wall detect that the internal soft tissues have filled the old exoskeleton and trigger production of a hormone from neurosecretory cells in the brain. This hormone acts upon the prothoracic gland, an endocrine gland in the prothorax, which in turn secretes the molting hormone, a steroid known as ecdysone. Molting hormone then acts on the epidermis, stimulating growth and cuticle formation. Metamorphosis likewise is controlled by a hormone. Throughout the young larval stages a small gland behind the brain, called the corpus allatum, secretes juvenile hormone (also known as neotenin). As long as this hormone is present in the blood the molting epidermal cells lay down a larval cuticle. In the last larval stage, juvenile hormone is no longer produced, and the insect undergoes metamorphosis into an adult. Among holometabolous insects the pupa develops in the presence of a very small amount of juvenile hormone.

Although a state of arrested development may occur during any stage, diapause occurs most commonly in pupae. In temperate latitudes many insects overwinter in the pupal stage (e.g., cocoons). The immediate cause of diapause, failure to secrete the growth and molting hormones, usually is induced by a decrease in daylength as summer wanes.

In addition to changes in form during development, many insects exhibit polymorphism as adults. For example, the worker and reproductive castes in ants and bees may be different, termites have a soldier caste as well as reproductives and persistent larvae, adult aphids (Homoptera) may be winged or wingless, and some butterflies show striking seasonal or sexual dimorphism. The general interpretation of all such differences is that, although the capacity to develop different forms is present in the genes of every member of a given species, particular lines of development are evoked by environmental stimuli. Hormones, including perhaps juvenile hormone, may be agents for the control of such changes.

Reproduction

The life of the adult insect is geared primarily to reproduction. Since reproduction is sexual in almost all insects, mating must be followed by impregnation of the female and fertilization of eggs. Usually the male seeks out the female. In butterflies in which vision is important, the colour of the female in flight can attract a male of the same species. In mayflies (Ephemeroptera) and certain midges (Diptera), males dance in swarms to provide a visual attraction for females. In certain beetles (e.g., fireflies and glowworms) parts of the fat body in the female have become modified to form a luminous organ that attracts the male. Male crickets and grasshoppers attract females by their chirping songs, and the male mosquito is lured by the sound emitted by the female in flight. The most important element in mating, however, is odour. Most female insects secrete odorous substances called pheromones that serve as specific attractants and excitants for males. The male likewise may produce scents that excite the female. Certain scales (androconia) on the wings of many male butterflies function in this way. Assembling scents, active in small quantities, are well known in female gypsy moths and silkworms as male attractants. The queen substance in the honeybee serves the same purpose.

Mating and egg production require appropriate temperatures and adequate nutrition. The need for protein is particularly important, and in insects such as Lepidoptera (butterflies and moths), which take only sugar and water in the adult stage, necessary protein is derived from larval reserves. Temperature and nutrition often influence hormone secretion. Juvenile hormone or hormones from the neurosecretory cells commonly are needed for egg production. In the absence of these hormones reproduction is arrested, and the insect enters a reproductive diapause. This phenomenon occurs in the potato beetle Leptinotarsa during the winter.

A few insects (e.g., the stick insect Carausius) rarely produce males, and the eggs develop without fertilization in a process known as parthenogenesis. During summer months in temperate latitudes, aphids occur only as parthenogenetic females in which embryos develop within the mother (viviparity). In certain gall midges (Diptera) oocytes start developing parthenogenetically in the ovaries of the larvae, and the young larvae escape by destroying the body of their mother in a process called paedogenesis.

Sensory perception and reception

Touch

Insects have an elaborate system of sense organs. Tactile hairs, concentrated on the antennae, palps, legs, and tarsi, cover the entire body surface. The hairs serve to inform the insect about its surroundings and its body position (a phenomenon known as proprioception). For example, contact between the hairs on the feet and the ground inhibits movement and may lead to a state of rest in some insects. Modified mechanical sense organs in the cuticle called campaniform organs detect bending strains in the integument. Such organs exist in the wings and enable the insect to control flight movements. Campaniform organs, well developed in small clublike halteres (the modified hind wings of dipterans), serve as strain gauges and enable the fly to control its equilibrium in flight.

Sound

Exceedingly sensitive organs called sensilla are concentrated in organs of hearing. These can be found on the bushy antennae of the male mosquito or tympanal organs in the front legs of crickets or in abdominal pits of grasshoppers and many moths. In moths these sensitive organs can perceive the high-pitched sounds emitted by bats as they hunt by echolocation. Insects complement organs of sound reception with sound-producing organs, which usually are (as in crickets) wing membranes that vibrate in response to movement of a stiff rod across a row of stout teeth. Sometimes (as in cicadas) a timbal (membrane) in the wall of the thorax is set in vibration by a rapidly contracting muscle attached to it.

Chemicals

Chemical perceptions by the thin-walled sensilla may be comparable to the human sense of taste or smell. Many insect chemoreceptors are specialized according to specific behaviour patterns. For example, although approximately equivalent to humans in the perception of flower odours and sugar sweetness, honeybees are exceedingly sensitive to the queen substance, which is scentless to humans. And male silkworm moths are excited by infinitesimal traces of the female sex pheromone, even in the presence of odours that are intensely strong to humans.

Sight

Although the insect eye provides less clarity than the human eye, insects can form adequate visual impressions of their surroundings. Insects have good colour vision, with colour perception extending (as in ants and bees) into the ultraviolet, although it often fails to extend into the deep red. Many flowers have patterns of ultraviolet reflection invisible to the human eye but visible to the insect eye.

Behaviour

Instincts

The insect orients itself by responding to the stimuli it receives. Formerly, insect behaviour was described as a series of movements in response to stimuli. That hypothesis has been supplanted by one that holds that the insect has a central nervous system with built-in patterns of behaviour or instincts that can be triggered by environmental stimuli. These responses are modified by the insect’s internal state, which has been affected by preceding stimuli. Patterns of behaviour range from comparatively simple reflex responses (e.g., the avoidance of adverse stimuli, the grasping of a rough surface on contact with the claws) to elaborate behavioral sequences (e.g., searching for mates, courtship, mating, and locating egg laying sites; hunting, capturing, and eating prey). The highest developments of behaviour, found in social insects such as the ants, bees, and termites, are based on the instinct principle.

An interesting example of a behavioral pattern is that found in the leaf-cutter bee Megachile. The female bee first locates a site for her nest in rotten wood and shapes the nest into a long tunnel. She then seeks out a preferred shrub from which pieces of leaves are gathered to build a cell. She first cuts a disc for a cell cap and then a series of oval pieces for the walls. After preparing the nest, she provisions it with a mixture of pollen and honey, lays an egg, and then closes the cell with more cut leaves. The leaf-cutter bee repeats this sequence until the nest is filled. Each act can be performed only in this set sequence. The insect does not stop to repair any damage to the nest but proceeds undeterred to the next step in her behavioral pattern.

Honeybee behaviours are more flexible than those of the leaf-cutter bee. Behavioral sequences of individuals are predictable, but the choice of acts or duties within the hive can be influenced by the needs of the colony. Honeybees exhibit capacity for learning (e.g., interpreting the waggle dance, learning flower colours), which is important in any insect that has to find its nest. Although these behaviours are necessary for both colony and food source location, learning capacity plays a relatively small part in the overall pattern of honeybee behaviour.

Experimental studies of details of behaviour have provided significant information about the properties of the sense organs. These studies also have provided information on the ability of insects to learn from their experience in the environment.

Insect societies

Both in complexity of behaviour and learning capacity, solitary wasps and bees are the equals of social wasps or honeybees. Social insects, however, have developed a division of labour in which the members must do the work required at the proper time. If the society is to succeed, its needs must be communicated to the individual members, and those individuals must act accordingly. These needs may be met by a temporary change in the behaviour of existing individuals, or they may result in developmental changes that vary the number of individuals in the various castes (e.g., new queens, males, workers, or soldiers). Commonly, both behavioral and developmental changes are initiated by pheromones, chemical messengers that convey information from one member of a colony to another.

Insect societies are gigantic families, with all individuals being the offspring of a single female. In the honeybee the single queen in the hive secretes a pheromone known as the queen substance (oxodecenoic acid), which is taken up by the workers and passed throughout the colony by food sharing. So long as the queen substance is present, all members are informed that the queen is healthy. If the workers are deprived of queen substance, they proceed at once to build queen cells and feed the young larvae with a special salivary secretion known as royal jelly that results in the production of new queens.

All termites and ants and some species of wasps and bees are the only insect groups containing truly social species. However, there are many other species that exhibit some lesser degree of interaction among individuals.

Ecology

Terrestrial insects

Insects feed on every sort of organic matter, and their methods of feeding and digestion have become modified accordingly. The major climatic hazards faced by terrestrial insects are temperature extremes and desiccation. Different species function best at various optimal temperatures. If conditions are too hot, an insect seeks out a cool, moist, and shady spot. If exposed to the sun on a hot day, an insect will position itself so as to present the smallest amount of body surface to the heat. If conditions are too cool, insects will remain in the sun to warm themselves. Many butterflies must spread their wings and expose the large surface to the sun like solar collectors to warm the flight muscles before they can fly. Many moths can raise their temperature by vibrating their wings or “shivering” before taking flight. The heat generated in this way is conserved by hairs or scales that maintain an insulating layer of air around the body. The optimum muscle temperature for flight is from 38° to 40° C (100° to 104° F).

In extremely cold weather the danger for insects is freezing, and insects that survive winters in cold latitudes are called cold hardy. A few insects (e.g., some caterpillars and aquatic midge larvae) tolerate ice formation in body fluids, although it is probable that the cell contents do not freeze. In most insects, however, cold hardiness means resistance to freezing. This resistance results partly from accumulation of large quantities of glycerol as an antifreeze and partly from physical changes in the blood that permit supercooling to temperatures far below the freezing point of water without the blood freezing.

Preventing water loss is another important aspect of life in terrestrial environments. All insects have a waxy (lipid) layer that coats the outer surface of the exoskeleton to prevent water loss from the body wall. In addition, most terrestrial insects also have adaptations to avoid water loss through respiration and waste elimination.

Aquatic insects

Major changes required for life in an aquatic habitat include modifications of the legs for swimming and adaptations for respiration. Most aquatic insects swim using the second or third (or both) pairs of legs. In some, the distal (away from the body) leg segments may simply be flattened and serve as oars. In others, there is a row of movable hairs on these segments that fold against the leg to offer less resistance during the forward stroke and then extend out, forming an oarlike surface during the power stroke. In some, like the water striders (Gerridae), long thin legs allow them to “walk” on the surface film of ponds and streams.

To breathe, some insects simply rise to the water surface and take atmospheric air into their tracheal systems. Mosquito larvae use only the last pair of abdominal spiracles, which open at the tip of a respiratory siphon. Water beetles (e.g., Dytiscus) have converted the space between the protective sheaths on the hind wings (elytra) and the abdomen into an air-storage chamber. Air-breathing insects can prolong the period of submergence by trapping air among their surface hairs. This air film acts as a physical gill and makes possible oxygen uptake from water. Other adaptations to an aquatic environment have occurred in larvae that obtain all their oxygen directly from the water. In midge larvae, abundant tracheae (breathing tubes) contact the entire thin cuticle. Caddisfly (Trichoptera) and mayfly (Ephemeroptera) larvae have tracheal gills on the abdomen or thorax. In dragonfly larvae, the gills are inside the rectum, and the water is pumped in and out through the anus, whereas damselflies have external rectal gills.

Protection from enemies

Insects may derive some protection from the horny or leathery cuticle but may also have various chemical defenses. Some caterpillars have special irritating hairs, which break up into barbed fragments that contain a poisonous substance that causes intense itching and serves as a protection against many birds.

Dermal glands of many insects discharge repellent or poisonous secretions over the cuticle, whereas others are protected by poisons that are present continuously in the blood and tissues. Such poisons often are derived from the plants on which the insects feed. In many hymenopterans (ants, bees, wasps) accessory glands in the female reproductive system have become modified to produce toxic proteins. These poisons, injected into the nervous system of the prey, paralyze it. In this state the prey serves as food for the wasp larva. Stings are also used by hymenopterans, including ants, wasps, and bees, for self-defense.

Concealment is an important protective device for insects. For some, this may be accomplished by simply hiding beneath stones or the bark of trees. However, many species rely on some forms of protective coloration. Protective coloration may take the form of camouflage (cryptic coloration) in which the insect blends into its background. The coloration of many insects copies a specific background with extraordinary detail. Stick insects (Carausius) can change their colour to match that of the background by moving pigment granules in their epidermal cells. Some caterpillars also have patterns that develop in response to a background, although these are irreversible. Insects such as caterpillars, which rely on cryptic coloration, often combine it with a rigid deathlike position.

Alternatively, insects that have well-developed chemical defenses generally show conspicuous warning (aposematic) coloration. Experiments have proved that predators such as birds quickly learn to associate such coloration “labels” with nauseous or dangerous prey. Finally, insects without nauseous qualities may gain protection by mimicry, that is, by developing a conspicuous colour pattern similar to that found in distasteful species (see also coloration; mimicry).

Population regulation

The factors that limit the numbers of insect species are complex. Experimental studies of a population of grain beetles in a container of wheat show that the complexities increase if a second species is added. With insects in natural habitats, competing not only with members of their own species but with numerous other species as well, the obstacles to survival become increasingly great. Competition among species is reduced to some extent by specialization of species to niches, or habitats, for which other insects do not compete.

Formerly, controversy arose over whether numbers were always density dependent (i.e., limited by the density of the species itself) or whether catastrophic actions, notably the vagaries of weather, were of prime importance. It has since become generally thought that the ultimate factor in the control of numbers is competition within the species for food and other needs. However, in many circumstances, before competition for food becomes significant, numbers are reduced by external factors. Competition within a species is often reduced by wholesale migration to new localities. Migration may occur by active flight or, as in aphids and locusts, largely directed by the wind. Another important factor in the regulation of populations is balanced polymorphism of species, in which the prevalence of individuals with given characteristics changes according to the action of natural selection as the state of the environment changes.

Form and function

External features

Cuticle

The insect is covered by the cuticle, a layer of inert material laid down by a single sheet of epidermal cells. It consists mainly of chitin, a carbohydrate also known as polyacetylglucosamine, and sclerotin, a hard substance composed of protein tanned by quinones. The cuticle, which has an outer layer of waterproofing wax to prevent loss of water by evaporation, also serves as the skeleton to which the muscles are attached. In insects such as caterpillars, in which the cuticle is soft and flexible, the skeleton is of the hydrostatic type. In this type, body fluid pressure, maintained by muscle tension beneath the body wall, provides the firmness necessary for the function of muscles involved in movement. In insects with hard bodies, the cuticle is made up of hardened areas called sclerites that are connected by flexible joints. At the back of the head and in the thorax, hardened ingrowths of the cuticle, known as apodemes, furnish a kind of internal skeleton for muscular attachment.

Insect colours depend partly on pigments incorporated in the cuticle. However, the most important pigments often occur in epidermal cells below the cuticle. In butterflies and moths, pigments may be deposited in flattened hairs, or scales, covering the wings. Some of the most brilliant insect colours are not the result of pigmentation but are physical interference colours produced by fine laminae (grooves or pits) in the surface of the wing scales or the cuticle itself.

Head

The ancestors of insects most likely had bodies consisting of many similar segments with only minor aggregation of the nervous system in the anterior (head) segment. These primitive insect ancestors probably looked something like modern centipedes, with a pair of appendages on each body segment but without a well-developed head. In present-day insects the primitive segments are grouped into three regions known as the head, thorax, and abdomen.

The first six primitive segments have fused to form the head, and the appendages of these segments have become modified into antennae that bear numerous sense organs and mouthparts that convey food to the mouth. Eyes also are prominent on the head. In most insects the mouthparts, adapted for chewing, consist of several parts; behind the upper lip or labrum is a pair of hard, toothed mandibles. These are followed by a pair of structures called first maxillae, each consisting of a bladelike lacinia, a hoodlike galea, and a segmented palp bearing sense organ. The paired second maxillae are partly fused in the midline to form the lower lip, or labium. Sometimes a median tonguelike structure, called the hypopharynx, arises from the floor of the mouth.

Insect mouthparts have been modified strikingly and reflect particular methods of feeding. The dipterans (true flies) provide instructive examples. In the primitive bloodsucking flies (e.g., the horsefly Tabanus) the mandibles and maxillae form serrated blades that cut through the skin and blood vessels of the host animal. The epipharynx and hypopharynx are elongated and grooved so that, when apposed, they form a tube for sucking blood. The tonguelike labium is used for imbibing exposed fluids. Dipteran mouthparts have evolved in two directions. In the mosquitoes (Culicidae) the mandibles, maxillae, epipharynx, and hypopharynx have become exceedingly slender stylets that form a fine bundle and are used for piercing skin and entering blood vessels. The labium, elongated and deeply grooved, serves only as a sheath for the stylet bundle. In the housefly Musca, however, mandibles and maxillae have been lost; the tonguelike labium alone remains and serves for feeding on exposed surfaces. Certain flies related to Musca have reacquired a capacity to suck blood; however, since they have lost both mandibles and maxillae, a new bloodsucking mechanism has developed. Labial teeth have evolved for cutting through the skin, and the labium itself is plunged into the tissues. The stable fly Stomoxys has an arrangement of this kind. In the tsetse fly Glossina, the labium has become a fine, needlelike structure normally protected by a sheath formed from the palps of the lost maxillae.

The insect thorax consists of three segments (called the prothorax, mesothorax, and metathorax), which may be fused but are usually recognizable. Each segment has four groups of hard plates (sclerites); the groups are the notum (upper), the pleura (sides), and the sternum (underside). Thoracic sclerites are located on a given segment by using an appropriate prefix (pro-, meso-, meta-); for example, the notum (upper sclerite) of the prothorax is the pronotum.

Each segment bears a pair of legs, and, in the mature insect, the mesothorax and metathorax typically carry a pair of wings. Each leg always consists of five parts: a coxa articulated to the thorax, a small trochanter, a femur, a tibia, and a tarsus with one to five segments. The tarsal segments often carry claws with adhesive pads between them (arolia or pulvilli); these enable the insect to hold onto smooth surfaces. The legs may be modified for leaping, burrowing, grasping prey, or swimming in various ways.

The wings at rest may be extended permanently on each side, as in some dragonflies (Odonata), or held erect above the body, as in mayflies (Ephemeroptera); in most insects, however, they are folded against the abdomen. The wing consists of cuticular sacs that bud out from the wall of the thorax; the sacs become flattened during development, and the two membranes, pressed together, are stiffened by thickenings of the cuticle that form cylindrical veins carrying tracheae, nerves, and circulating blood to all parts of the wing. Wings utilized for flight commonly are made of thin membranous cuticle. In some insects, notably beetles (Coleoptera), the wings of the middle segment of the thorax have become thick and horny and serve as protective sheaths (elytra) of the membranous hindwings.

The locomotion of insects is effected by muscles acting on the external skeleton. In leaping insects (e.g., grasshoppers, fleas) the force of muscle contraction is used to compress a pad of an elastic protein, resilin; when the catch mechanism is released, the stored energy in the protein molecule is used to project the insect into the air. Insect flight is achieved by flapping the wings; during these movements the wing blade, twisted as it passes from elevation to depression, produces the same effect as the rotating propeller of an aircraft. Muscles capable of changing this inclination control the direction of flight. The chief flight muscles control flight in one of two ways: in dragonflies, directly on a lever at the base of each wing; but, in most insects, indirectly by deforming the shape of the thorax. The longitudinal muscles of the thorax depress the wings that are articulated with it; the vertical muscles elevate them.

In butterflies, the number of wing beats per second may be as low as 8 to 12, while the rate in mosquitoes may exceed 600. These rates can exceed the frequency of contraction and relaxation of muscles responding to nerves because the muscles, after they have begun contracting and relaxing, respond to the alternating elastic tension in the thoracic wall, where the frequency is determined by the natural periodic oscillation of the thorax. The flight of insects, despite their small size, conforms to the aerodynamic laws that regulate the flight of aircraft.

Abdomen

The abdomen consists of a maximum of 11 segments, although this number commonly is reduced by fusion. Appendages are usually absent except in caterpillars, which use up to five pairs of abdominal prolegs in walking, and in adult insects where the appendages at the hind end have become transformed into external genitalia. In the male these genitalia are paired claspers used to hold the female; in the female, three pairs of valvulae are used to manipulate eggs during oviposition. In some insects, notably crickets and cockroaches, two feelers, or cerci, at the hind end of the abdomen bear sense organs.

Internal features

Digestive system

The nutritive requirements of insects are much the same as those of mammals—water, inorganic ions, and essential amino acids (i.e., those that cannot be synthesized by the animal). The requirements for preformed fat and carbohydrate vary with the species. Although vitamins of the B group are needed by insects, neither vitamins A nor D are required, and many insects can synthesize ascorbic acid (vitamin C). On the other hand, insects cannot synthesize adequate quantities of cholesterol; thus, in effect, cholesterol can be defined as a vitamin for insects.

Insects that feed solely on some restricted diet (e.g., sterile blood, plant juices, refined flour) have special cells termed mycetocytes that harbour symbiotic micro-organisms; these organisms, transmitted through the egg to the next generation, benefit their host by furnishing it with an internal source of vitamins and perhaps other essential nutrients. If the symbiotic micro-organisms are removed experimentally, an insect fails to grow if not provided with a diet rich in vitamins.

The digestive system consists of a foregut formed from the mouth region (stomodaeum), a hindgut formed similarly from the anal region (proctodaeum), and a midgut (mesenteron). The foregut and hindgut are lined by cuticle continuous with that on the body surface. The mouth is followed by the muscular pharynx, which functions in sucking and swallowing, and the esophagus, which may be enlarged to form a crop. The crop discharges into the midgut, sometimes, as in cockroaches, by way of a muscular gizzard or proventriculus. The termination of the midgut is marked by the attachment of the malpighian tubules, the chief organs of excretion. The hindgut commonly consists of a narrow ileum followed by a larger and often thick-walled rectum, which discharges at the anus.

Digestive enzymes, secreted not only by the salivary glands but also by the cells of the midgut and its diverticula, vary with the diet of the insect. The most important enzyme secreted by the salivary glands is amylase; the midgut secretes several enzymes including protease, lipase, amylase, and invertase. The products of digestion are absorbed chiefly in the midgut.

The hindgut receives food residues from the midgut as well as waste products from the malpighian tubules. The end products of nitrogen metabolism are uric acid, small amounts of amino acids, and urea; in aquatic insects, ammonium salts may be a major form for nitrogen excretion. In the rectum, the epithelial cells lining the gut wall often are enlarged, particularly in restricted areas where they form rectal glands. The epithelial cells of these glands are supplied richly with tracheae and function in the reabsorption of water and ions. The rectal contents of insects that inhabit dry environments commonly are reduced to dry fecal pellets prior to discharge. In many insects, particularly those which feed on relatively dry foods (e.g., beetles infesting stored grain), the upper segments of the malpighian tubules are bound by a sheath to the rectal surface and form a cryptonephridial system that serves to increase the capacity of the rectum for reabsorbing water and salts. The products of digestion, discharged into the hemocoele, or general body cavity, are transported by the circulatory fluid, or hemolymph, to the organs.

Circulatory system

The circulatory system is an open one, with most of the body fluid, or hemolymph, occupying cavities of the body and its appendages. The one closed organ, called the dorsal vessel, extends from the hind end through the thorax to the head; it is a continuous tube with two regions, the heart or pumping organ, which is restricted to the abdomen, and the aorta, or conducting vessel, which extends forward through the thorax to the head. Hemolymph, pumped forward from the hind end and the sides of the body along the dorsal vessel, passes through a series of valved chambers, each containing a pair of lateral openings called ostia, to the aorta and is discharged in the front of the head. Accessory pumps carry the hemolymph through the wings and along the antennae and legs before it flows backward again to the abdomen.

The circulating hemolymph, or blood, is not important in respiration but functions in transporting nutrients to all parts of the body and metabolic waste products from the organs to the malpighian tubules for excretion. It contains free cells called hemocytes, most of which are phagocytes that help to protect the insect by devouring micro-organisms. An important tissue bathed by the hemolymph is the fat body, the main organ of intermediary metabolism. It serves for the storage of fat, glycogen, and protein, particularly during metamorphosis. These materials are set free as required by the tissues for energy production or for growth and reproduction.

Respiratory system

The respiratory system consists of air-filled tubes or tracheae, which open at the surface of the thorax and abdomen through paired spiracles. The muscular valves of the spiracles, closed most of the time, open only to allow the uptake of oxygen and the escape of carbon dioxide. The tracheal tubes are continuous with the cuticle of the body surface. The tracheae are stiffened by spiral thickenings or threadlike ridges called taenidia, which branch repeatedly, becoming reduced in cross section and ending in fine thin-walled tracheoles less than one micron in diameter. The tracheoles insinuate themselves between cells, sometimes appearing to penetrate into them, and push deeply into the plasma membrane.

Although movements of oxygen and carbon dioxide occur solely by gaseous diffusion in sedentary insects, the system is ventilated mechanically in active species. Pumping movements of the abdomen provide the force necessary to drive out streams of air at some spiracles and suck them in at others. The taenidia keep the tracheae distended, thus allowing free passage of air. In addition, the most active insects have large thin-walled dilatations of the tracheae called air sacs, which serve to increase the volume of air displaced during respiratory movements. Both lack of oxygen and accumulation of carbon dioxide provide stimuli to nerve centres that induce increased respiration during muscular activity.

Reproductive system

The reproductive system consists of the sex glands, or gonads (male testes and female ovaries), the ducts through which the sexual products are carried to the exterior, and the accessory glands. The two testes are made up of a variable number of follicles in which the spermatocytes mature and form packets of elongated spermatozoa. Spermatozoa, liberated in bundles with heads held in a cap of gelatinous material, accumulate in the vesicula seminalis, a dilated section of the male sexual duct (vas deferens).

Each of the two ovaries consists of a number of ovarioles. The ovarioles converge upon the two oviducts, and the oviducts unite to form a common oviduct down which the ripe eggs are discharged. Each ovariole consists of a germarium and a series of ovarial follicles. The germarium is a mass of undifferentiated cells that form oocytes, nurse cells, and follicular cells. The nurse cells provide nourishment for the oocytes during the early stages of their growth; follicular cells, which invest the enlarging oocyte as a continuous epithelium, provide the materials for yolk formation and, in the final stages, lay down the eggshell or chorion. The ovarial follicles increase progressively in size as the oocytes grow to form ripe eggs.

During copulation, bundles of spermatozoa are sometimes introduced directly into the female vagina by means of the male copulatory organ, or aedeagus. Secretions from the accessory glands of the female activate the sperm, the sperm bundles disperse, and the free spermatozoa make their way up to the receptaculum seminis, or spermatheca, where they are stored, ready to fertilize the eggs. In most insects, the male accessory glands secrete materials that form a tough capsule, or spermatophore; spermatozoa are encased in this spermatophore, which is inserted into the entrance of the vagina. The spermatophore walls commonly contain a gelatinous substance that swells upon exposure to secretions of the female and forces out the spermatozoa. The vagina serves both for receiving sperm and for laying eggs.

The terminal segments of the abdomen of females sometimes are modified to form an ovipositor used for depositing eggs. In butterflies and moths (Lepidoptera) a second copulatory canal independent of the vagina has been evolved, so that the sperm enter by one route, and the eggs are deposited by another.

The eggshell, or chorion, commonly provided with an air-filled meshwork, provides for respiration of the developing embryo. The chorion is also pierced by micropyles, fine canals that permit entry of one or more spermatozoa for fertilization. As the egg passes down the oviduct before egg laying, the micropyles come to lie opposite the duct of the spermatheca; at this stage fertilization occurs. Eggs must be waterproof to prevent desiccation; each egg has a layer of waterproofing wax, sometimes over the entire shell surface, more often lining the inside.

Nervous system

The central nervous system consists of a series of ganglia that supply nerves to successive segments of the body. The three main ganglia in the head (protocerebrum, deutocerebrum, and tritocerebrum) commonly are fused to form the brain, or supraesophageal ganglion. The rest of the ganglionic chain lies below the alimentary canal against the ventral body surface. The brain is joined by paired connectives to the subesophageal ganglion, which is linked in turn by paired connectives to the three thoracic and eight abdominal ganglia (numbered according to segment). In most insects the number of separate ganglia has been reduced by fusion. The last abdominal ganglion always serves several segments. In homopterans and heteropterans all the abdominal ganglia usually fuse with mesothoracic and metathoracic ganglia; and in the larvae of higher flies (Cyclorrhapha), the ganglia of the brain, thorax, and abdomen form one mass.

Each ganglion is made up of nerve-cell bodies that lie on the periphery and a mass of nerve fibres, the neuropile, that occupies the centre. There are two types of nerve cells, motor neurons and association neurons. Motor neurons have main processes, or axons, that extend from the ganglia to contractile muscles, and minor processes, or dendrites, that connect with the neuropile. Association neurons, usually smaller than motor neurons, are linked with other parts of the nervous system by way of the neuropile.

Cell bodies of the sense organs, called sensory neurons, lie at the periphery of the body just below the cuticle. Sensory neurons occur as single cells or small clusters of cells; the distal process, or dendrite, of each cell extends to a cuticular sense organ (sensillum). The sensilla are usually small hairs modified for perception of specific stimuli (e.g., touch, smell, taste, heat, cold); each sensillum consists of one sense cell and one nerve fibre. Although these small sense organs occur all over the body, they are particularly abundant in antennae, palps, and cerci. The sense cell of each sensillum gives off a proximal process, or sensory axon, which runs inward to the central nervous system, where it enters the neuropile and makes contact with the endings of association neurons. Bundles of both sensory axons and motor axons, which are enclosed in protective membranous sheaths, constitute the nerves.

Tactile hairs may be sensitive enough to perceive air vibrations and thus serve as organs for sound reception. Tympanal organs (eardrums) are present in certain butterflies and grasshoppers. Mechanical sensilla (chordotonal organs) below the surface of the cuticle serve for perception of internal strains and body movements.

Eyes

The eyes are of two kinds, simple eyes, or ocelli, and compound eyes. In the adults of higher insects both types are present. The visual sense cells are derived from the epidermis, as are those of other sense organs, and are connected to the optic ganglia (a part of the brain) by sensory axons. Each visual sense cell has a zone at its surface, which, on exposure to light, gives rise to chemical products that stimulate the sense cell, called the retinula cell, and initiate the nerve impulse in the sensory axon. The light-receptive zone, or rhabdom, of the retinula cell commonly has a rodlike form; because it lies perpendicular to the surface, light passes lengthwise along it. In the simple eyes (ocelli) a lens-shaped area of cuticle lies over the group of retinula cells that form the retina. Since the optical structure is primitive, the visual image received is crude; ocelli can perceive only light, darkness, and movement.

The compound eye, made up of a number of facets, resembles a honeycomb; each facet overlies a group of six or seven retinal cells that surround the rhabdom. Each of the retinal units below a single facet is termed an ommatidium. The number of facets varies. For example, there are only a few dozen facets in the eye of the primitive apterygote Collembola, while the eye of the housefly Musca has some 4,000, and the highly developed eye of the dragonfly may contain up to 28,000.

During light reception, rays from a small area of the field of view fall on a single facet and are concentrated upon the rhabdom of the retinula cells below. Since each point of light differs in brightness, all the ommatidia that form the retina receive a crude mosaic of the field of view. Unlike the image in a camera or in human eyes, the mosaic image in the compound eye is not inverted but erect. The fineness of the mosaic and, therefore, the degree of resolution improves with increasing numbers of facets. It is estimated that the eye of the honeybee has visual acuity equal to 1 percent that in man.

Each ommatidium commonly is shielded by a curtain of pigmented cells that prevent the spread of light to neighbouring ommatidia. This is termed an apposition eye. In the eyes of insects that fly at night or in twilight, however, the pigment can be withdrawn so that light received from neighbouring facets overlaps to some extent. This is termed a superposition eye. The image formed is brighter but not as sharp as that formed by the apposition eye. In addition to perceiving brightness, the eyes of insects can perceive colour as well as some other properties of light.

Evolution and paleontology

Origin of insects

The most primitive insects known are found as fossils in rocks of the Middle Devonian Period and lived about 350,000,000 years ago. The bodies of those insects were divided then, as now, into a head bearing one pair of antennae, a thorax with three pairs of legs, and a segmented abdomen. Those insects originated with the terrestrial branch of the phylum Arthropoda. The Arthropoda, whose origin is thus far unknown, probably arose in Precambrian times, perhaps as much as 1,000,000,000 years ago. Some arthropods colonized the open sea and have become the present-day class Crustacea (crabs, shrimps) and the now-extinct Trilobita. Other arthropods colonized the land. This terrestrial line persists chiefly as the classes Onychophora, Arachnida (spiders, scorpions, ticks), the myriapods (consisting of Diplopoda [millipedes], Pauropoda, Symphyla, and Chilopoda, or centipedes), and finally the class Insecta.

The most primitive insects today are found among the wingless (apterous) hexapods; sometimes known collectively as apterygotes, they include proturans, thysanurans, diplurans, and collembolans. It is agreed generally that insects are related most closely to the myriapod group, among which the Symphyla exhibit most of the essential features required for the ancestral insect form (i.e., a Y-shaped epicranial suture, two pairs of maxillae, a single pair of antennae, styli and sacs on the abdominal segments, cerci, and malpighian tubules). There is, therefore, general agreement that the insects probably arose from an early symphylan-like form.

Insect fossil record

The insect fossil record has many gaps. Among the primitive apterygotes, only the collembolans (springtails) have been found as fossils in the Devonian Period. Ten insect orders are known as fossils, mostly of Late Carboniferous and Permian times. No fossils have yet been found from the Late Devonian or Early Carboniferous periods, when the key characters of present-day insects are believed to have evolved; thus, early evolution must be inferred from the morphology of extant insects.

It has become evident that insect evolution, like that of other animals, was far more active at some periods than at others. There have been geological epochs of “explosive” evolution during which many new forms have appeared. Those epochs may have followed some modification or innovation in body function, or new developments favoured by climatic changes or evolutionary advances of other animals and plants. During those periods of evolutionary change, new methods of feeding and living led to diversity of insect mouthparts and limbs, the origin of metamorphosis, and other changes.

Insect phylogeny

Figure 1 is a simplified family tree of the presumed evolutionary history of winged insects (Pterygota) throughout the geological periods from the Devonian to the Recent. The apterygotes, which are regarded as survivors of primitive insect stock, are omitted from the family tree. Dark lines indicate the periods during which the various orders have been found as fossils. Some lines stop at the names of orders now extinct and known only as fossils. Light lines indicate the hypothetical origin of various orders. Many insect types, traces of which have not yet been discovered, must have been produced during the explosive periods of evolution in Carboniferous and Permian times.

The primitive wingless insects (Figure 1) gave rise to a paleopterous stock. Descendants of this stock included ancient fossil types that flourished in Permian times, such as the giant dragonflies or Protodonata (some of which had a wing span of more than half a metre) and dragonflies and damselflies (Odonata) and mayflies (Ephemeroptera), both of which have persisted with little change to the present. The primitive insect stock also gave rise to a neopterous stock, believed to include the progenitors of the remaining insect orders. The Orthoptera (grasshoppers) and the Plecoptera (stoneflies) have been found as fossils even in late Carboniferous times. The Isoptera (termites, sometimes placed in the order Blattodea), Embioptera (webspinners), and Dermaptera (earwigs), though doubtless of ancient origin, have not been found yet as fossils dated earlier than the Mesozoic Era.

The evolutionary radiation (Figure 1), believed to have given rise to the orders listed above in the Middle Carboniferous Period, is thought to have produced also a paraneopterous stock, which formed the base for a new evolutionary radiation during the Permian Period. Present-day derivatives of this stock evolved into the Psocoptera (psocids), Mallophaga (chewing lice), Anoplura or Siphunculata (sucking lice), Thysanoptera (thrips), Heteroptera (true bugs), and Homoptera (e.g., aphids).

Several phylogenetic lines (Figure 1) are exopterygote (i.e., insects with simple metamorphosis) some of which, such as Mallophaga and Anoplura, are secondarily wingless. The remaining orders are endopterygote (insects with complete metamorphosis). They are shown in Figure 1 as derivatives of an oligoneopterous stock, which gave rise to Neuroptera (lacewings), Hymenoptera (ants, wasps, and bees), and Coleoptera (beetles) in the Early Permian Period; the early ancestry of these orders is obscure, however, and the earliest fossils closely resemble present-day forms. One line from the evolutionary radiation (Figure 1) at the beginning of the Permian gave rise to a mecopteroid stock, and there is good evidence that a sub-radiation of these mecopteroid orders (sometimes called the panorpoid complex) provided the origin for the present Mecoptera (scorpionflies), Diptera (true flies), Siphonaptera (fleas), Trichoptera (caddisflies), and Lepidoptera (butterflies and moths).

Evolution

Wings and flight

Insect wings develop as paired outgrowths from the thorax, stiffened by ribs, or veins, in which run tracheae. These tracheae follow a consistent pattern throughout the Pterygota, and their specific modifications (known as venation) are important in classification and in estimations of the degree of relationship between groups. The basic consistency of venation suggests that wings have been evolved only once among the insects, that is, all the Pterygota (as shown in Figure 1) arose from a single stem. By the time (toward the end of the Carboniferous) fossil insects are found, wings are developed fully. In the Paleoptera (Figure 1) the wings are held aloft above the back, as in mayflies, or held extended permanently on each side of the body, as in dragonflies. Throughout the Neoptera there is a wing-flexing mechanism (secondarily lost in butterflies) that enables the wings to be folded back to rest on the surface of the abdomen.

Winged insects must have made their appearance very early in the Carboniferous, more than 300,000,000 years ago; but there is no fossil evidence to show the way they evolved. One hypothesis is that wings arose as fixed planes extending sideways from the thorax and that these planes were used, perhaps in some large leaping insect, for gliding. Later muscles developed, first to control inclination and then to move the wings in flapping flight. Another hypothesis is that wings may have originated from large thoracic tracheal gills, similar to the movable tracheal gills along the abdomen of some mayfly larvae. Such outgrowths could have been useful to insects exposed by the drying up of a temporary aquatic habitat and might have carried them in rain-bearing winds to a new watery home. It is likely that the most primitive symphylan-like insects were terrestrial; throughout insect evolution, however, independent adaptations to aquatic habitats have occurred. Usually the pattern is one in which the adults leave the water and disperse. Many pterygote insects have become secondarily wingless, sometimes as single species or groups of species within large orders, sometimes as entire orders (the parasitic lice, Mallophaga and Anoplura, and the fleas, Siphonaptera).

Metamorphosis

It generally is agreed that insect metamorphosis evolved as adult insects gradually adopted different modes of life from those of larvae. The characters of larva and adult became genetically independent; in response to natural selection, therefore, each was able to evolve independently of the other. Mouthparts, limbs, and other morphological features were modified in different directions and in higher groups. Where these differences were extreme, an intermediate pupal stage evolved to bridge the morphological gap between larva and adult. It seems quite probable that the development of metamorphosis occurred more than once during the evolution of insects.

Feeding methods

Insects did not evolve in a constant environment. Throughout geological time there were prodigious changes in climate; in addition, evolution was continuous among all other animals and plants. Geologically the selection pressures among insects were changing continuously. At the end of the Mesozoic Era the first flowering plants appeared. Insect evolution has paralleled that of the flowering plants; they have evolved together. As Lepidoptera (butterflies and moths), Hymenoptera (ants, bees, and wasps), Diptera (true flies), and Coleoptera (beetles) began to feed upon flowers, nectar, or pollen, flowering plants came to rely more and more upon insects—rather than upon the wind—for transferring their pollen. Flowers evolved nectaries, scents, and conspicuous colours as attractants for those insects that could effect cross-pollination. Insects likewise evolved appropriate mouthpart modifications for extracting nectar from flowers.

During the Mesozoic warm-blooded animals (mammals and birds) first appeared; by the dawn of the Tertiary Period, they had become predominant among the earth’s large animals. The warm fermenting excrement and the decaying dead bodies of mammals furnished excellent nutrient media for many insect larvae, notably among the Diptera and Coleoptera. The adults in both groups found their nourishment in flowers. Some heteropterans (true bugs) and dipterans pierce the skin of birds and mammals and feed on their blood. The Anoplura (sucking lice) and the Siphonaptera (fleas) have become so specialized for this type of parasitic existence that their relationships to other insects are not yet known with certainty.

Continuing evolution

Evolution is occurring among present-day insects. They exhibit a balanced genetic polymorphism; in other words, in response to small environmental changes, one genetic form, more successful than another, will become more plentiful. Sometimes there is no visible difference between these forms, the advantage presumably lying in some physiological change. It is advantageous for a species to have a gene pool from which favourable characters can be selected so that the species can respond to environmental changes. Changes within a species may occur progressively over a large geographical area. Such a progressive genetic change is termed a cline; in some cases insects at the extremes of the cline are so unlike that they are taken as separate species and may be infertile when crossed.

One well-known example of evolution in action among insects is industrial melanism (accumulation of the black pigment melanin); many butterflies inhabiting industrial areas have become almost black during the past century; black forms are more tolerant of pollution and less conspicuous to predators. Another example of this cline type of evolution is the development of insect strains resistant to an insecticide that has been applied heavily in an area for several years. In many parts of the world houseflies have become highly resistant to DDT.